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Revista Mexicana de Biodiversidad 83: 1036-1044, 2012DOI:
10.7550/rmb.28124
Recibido: 20 octubre 2011; aceptado: 02 junio 2012
Stem and root anatomy of two species of Echinopsis
(Trichocereeae: Cactaceae)
Anatoma de la raz y del tallo de dos especies de Echinopsis
(Trichocereeae: Cactaceae)
Joelma dos Santos Garcia1, Edna Scremin-Dias1 and Patricia
Soffiatti2
1Universidade Federal de Mato Grosso do Sul, CCBS, Departamento
de Biologia, Programa de Ps Graduao em Biologia Vegetal Cidade
Universitria, S/N, Caixa Postal 549, CEP 79.070.900 Campo Grande,
MS, Brasil.2Universidade Federal do Paran, SCB, Departamento de
Botnica, Programa de Ps-Graduao em Botnica, Caixa Postal 19031, CEP
81.531.990 Curitiba, PR, Brasil.
[email protected]
Abstract. This study characterizes and compares the stem and
root anatomy of Echinopsis calochlora and E. rhodotricha
(Cactaceae) occurring in the Central-Western Region of Brazil, in
Mato Grosso do Sul State. Three individuals of each species were
collected, fixed, stored and prepared following usual anatomy
techniques, for subsequent observation in light and scanning
electronic microscopy. Echinopsis calochlora revealed uniseriated
epidermis, while E. rhodotricha had patches of bisseriated
epidermis; all species showed thick cuticle, parallelocytic stomata
at the epidermis level, and a well-developed hypodermis. Cortical
and medullary bundles are present in the studied species, as well
as mucilage cells in the cortex region. The secondary phloem is
composed by sieve tube elements, companion cells, axial and radial
parenchyma. Sclereids were found at the outer regions of phloem in
the roots. The secondary xylem is non fibrous in the stems of E.
calochlora, and fibrous in the stems of E. rhodotricha and in the
roots of both species. Many of these characteristics are commonly
found in Cactaceae, and represent important adaptations for
survival in xeric environments.
Key words: cortical bundles, epidermis, non fibrous wood, wood
anatomy.
Resumen. Este estudio est enfocado a caracterizar y comparar la
anatoma de tallos y races de Echinopsis calochlora y E. rhodotricha
(Cactaceae) que habitan en la regin centro-oeste de Brasil, en el
Estado de Mato Grosso do Sul. Se recolectaron 3 individuos de cada
especie, los cuales fueron fijados, almacenados y preparados
siguiendo las tcnicas comunes de anatoma, para observarlos en
microscopa de luz y electrnica de barrido. Echinopsis calochlora
mostr epidermis uniseriada, mientras que la de E. rhodotricha fue
biseriada; todas las especies presentaron cutcula gruesa, estomas
paralelocticos a nivel de la epidermis y una hipodermis bien
desarrollada. Se presentaron haces vasculares corticales y
medulares en las especies estudiadas, as como clulas mucilaginosas
en la regin cortical. El floema secundario est compuesto de
elementos de tubo criboso, clulas acompaantes y parnquima axial y
radial. Se encontraron esclereidas en las regiones externas del
floema en las races. El xilema secundario es no-fibroso en los
tallos de E. calochlora y fibroso en los de E. rhodotricha y en las
races de ambas especies. Muchas de estas caractersticas estn
presentes comnmente en especies de Cactaceae, lo que representa
adaptaciones importantes para la supervivencia en ambientes
xricos.
Palabras clave: haces corticales, epidermis, madera no-fibrosa,
anatoma de la madera.
Gibson and Nobel, 1986; Soffiatti and Angyalossy, 2003, 2005,
2007; Terrazas and Arias, 2003; Arruda et al., 2005), considering
the representativeness of the family, and the problems in
circumscribing groups, especially within tribes, as well as
enabling a better understanding of the group evolution (Terrazas
and Arias, 2003) and their adaptations for survival in arid
environments (Mauseth, 2006).
Eggli (2002) mentioned the lack of information on Cactaceae in
Brazil in the Brazilian southwestern region, elaborating a
checklist of 33 native species for the states of Mato Grosso and
Mato Grosso do Sul. From those, 27
Introduction
The subfamily Cactoideae (Cactaceae) comprises the largest
number of species, is the most diversified in terms of life-forms
and habit (Taylor, 2000; Terrazas and Arias, 2003), and it is
divided in 9 tribes, in which Cereeae, Rhipsalideae, Trichocereeae,
Echinocereeae, and Hylocereeae occur in Brazil (Taylor and Zappi,
2004).
The importance of anatomical studies in Cactaceae are undeniable
(Gibson and Horak, 1978; Gasson, 1981;
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1037Revista Mexicana de Biodiversidad 83: 1036-1044, 2012DOI:
10.7550/rmb.28124
species belong to Cactoideae, and 6 genera were recorded for
tribe Trichocereeae: Arthrocereus (A. Berger) Backeb. et F.M.
Knuth, Cleistocactus Lem., Discocactus Pfeiff., Echinopsis Zucc.,
Gymnocalycium Pfeiffer in Pfeiffer et Otto, and Harrisia Britton,
all being very poorly studied, especially in the state of Mato
Grosso do Sul.
Echinopsis is one of the largest genus of Trichocereeae,
composed by 128 species distributed in several regions of South
America, and due to the large number of species, it shows a very
controversial circumscription (Anderson, 2001). Echinopsis
calochlora K. Schum. is endemic to the high altitudes of the Macio
do Urucum (Eggli, 2002), in western Mato Grosso do Sul, and was
recently included in the Brazilian list of threatened species
(IBAMA, 2008), because its habitat has been intensively exploited,
for iron and manganese mining (Silva et al., 2000). Echinopsis
rhodotricha K. Schum. has a broader distribution, occurring Brazil,
Paraguay and Argentina (Anderson, 2001). This work describes
comparatively the anatomy of stem and root of E. calochlora and E.
rhodotricha.
Materials and methods
Individuals were collected in rocky outcrops and sandy-clay
formations from natural populations as follows: E. calochlora at
Corumb County, in Pantanal (voucher number CGMS 17536; J. S. Garcia
54), and E. rhodotricha, at Porto Murtinho, in Chaco (voucher
number CGMS 17590; J. S. Garcia 61). Vouchers were deposited at the
Herbarium - CGMS/UFMS, Federal University of Mato Grosso do
Sul.
For the qualitative anatomical study of the stem, 3 individuals
of each species were sectioned at the basal, medium, and apical
regions. For the root, only the very base was used. Samples were
fixed with FAA 70 for 72 hours, rinsed in water and stored in
ethanol 70% (Jensen, 1962). Paradermal sections were taken at the
median region of the stem. Transverse and longitudinal sections of
stems and roots were made by hand. Samples were also embedded in
polyethylene glycol (PEG) 1500 (Richter, 1985) and transverse and
longitudinal sections were made in rotative microtome. Root samples
of E. rhodotricha were sectioned in a sliding microtome. Sections
were stained with astrablue and safranin and mounted in glycerine
50%. Macerations were prepared using a modified Franklin method
(Franklin, 1945), stained with safranin and mounted in
glycerin.
The following hystochemical tests were carried out: floroglucyn
(Johansen, 1940) to detect lignified secondary walls; lugol
(Johansen, 1940) to detect starch; for mucilage we followed Richter
(1977). The analysis was carried out in a Leica DMLB light
microscope, connected to an image capture system, and a digital
camera DC 300F.
Paradermal and transversal sections of the dermal system of the
stem median region were prepared for scanning electron microscopy
(SEM) analysis. Samples were dehydrated, critical-point dried,
mounted on aluminum stubs, and coated with gold-palladium in a
sputter system. SEM analysis and electron micrographs were made
with a JEOL JSM 5800 scanning electron microscope.
Results
Stem. Uniseriate epidermis (Fig. 1) occurred in
photosynthesizing regions of the stem, as well as in apical and
median regions. It was composed by square to rectangular cells in
cross sections, with thick cuticle. Echinopsis rhodotricha showed
patches of bisseriate epidermis in the median region (Fig. 2),
formed by periclinal divisions of epidermal cells, rarely observed
in E. calochlora.
Parallelocytic stomata were found in both species at the same
level of epidermal cells in E. calochlora (Fig. 1) while E.
rhodotricha had sunken stomata (Fig. 2). Substomatal chambers
crossed the hypodermis (Figs. 1, 2).
The phellogen was originated in the epidermis. Peridermis
covered the non photosynthesizing portions of the stem (Figs. 3,
4); composed of pheloderm formed by rectangular cells in cross
section, with primary walls. Cork had layers of cells with
suberized walls that alternate with layers of cells with lignified
walls, which are very numerous in E. rhodotricha (Fig. 4).
We observed a collenchymatic hypodermis in the stem (Figs. 1, 2,
5), composed of cells with irregularly thickened primary walls,
connecting through ramified channels.
Cortex was divided in an outer region forming a palisade (Fig.
5), and an inner region composed of large parenchyma cells and
vascular bundles (Fig. 5), constituted by phloem and xylem,
arranged collaterally. Druses were rare in the cortex. Large
mucilage cells occurred throughout the whole cortex (Fig. 6) and
pith. Vascular bundles occurred also in the pith (Fig. 7).
At the base of stem, primary phloem was collapsed and in E.
rhodotricha caps of sclereids were seen outside secondary phloem
(Fig. 8). Secondary phloem was composed of narrow sieve tube
elements and companion cells, with simple sieve plate, transverse
to slight oblique, axial and radial parenchyma.
Both species had non fibrous wood, mainly composed of wide band
tracheids and a few vessel elements, surrounded by axial and radial
unlignified parenchyma cells (Figs. 9-11). Wide band tracheids
presented ring, helical or mixed wall thickenings (Fig. 11). Vessel
elements had simple perforation plates. Both species had a storied
structure in the non fibrous wood (Fig. 11), being
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1038 Garcia et al.- Anatomy of two species of Echinopsis
Figures 1-6. Stem cross sections. 1-2, epidermis covered by a
thick cuticle (arrow). Stomata chamber crossed hypodermis. 1,
Echinopsis calochlora. Uniseriate epidermis. Stomata at the same
level as ordinary epidermal cells. 2, E. rhodotricha. Scanning
electron micrograph (SEM). Bisseriate epidermis and sunken stomata.
3-4, peridermis with pheloderm composed of layers of parenchyma
cells; cork composed of layers of cells with suberized walls
alternating with layers of cells with lignified walls (*). 3, E.
calochlora. 4-6, E. rhodotricha. 5, organization of cortex:
hypodermis followed by layers of palisade parenchyma and inner
isodiametric parenchyma cells. Cortical bundles (arrows). 6,
cortex. Numerous scattered mucilage cells. Scale bars: 1= 100 m; 2=
5 m; 3-4= 200 m; 5= 500 m; 6= 250 m. Cm- mucilage cell; Ep-
epidermis; Hp- hypodermis; Pp- palisade parenchyma; St- stomata;
Su- suber.
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1039Revista Mexicana de Biodiversidad 83: 1036-1044, 2012DOI:
10.7550/rmb.28124
Figures 7-12. 7-10, 12, cross sections. 11, tangential section.
7, Echinopsis calochlora. Pith with medullary bundle. 8, E.
rhodotricha. Vascular system. Phloem. Caps of sclereids in the
periphery of secondary phloem. 9, E. calochlora. Non fibrous wood
mainly composed of wide band tracheids. 10, E. rhodotricha. Fibrous
and non fibrous wood. 11, E. calochlora. Non fibrous wood composed
of wide band tracheids with helical thickenings. 12, E.
rhodotricha. Fibrous wood composed of vessel elements, fibres and
scanty paratracheal parenchyma (part of a ray is seen on the left).
Scale bars: 7, 9-12= 200 m; 8= 400 m. FW- fibrous wood; NFW- non
fibrous wood.
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1040 Garcia et al.- Anatomy of two species of Echinopsis
most pronounced in E. calochlora. Fibrous wood occurred only in
E. rhodotricha, outside non fibrous wood (Figs.10, 12-15), diffuse
porous, with wide band tracheids absent, composed of vessels
predominantly solitary and multiples of 2 to 3, and less frequent,
of 4 to 5; vessel elements walls had helical to pitted secondary
walls; intervessel pits were bordered; vessel-parenchyma pits
simple to half-bordered, scalariform, opposite to alternate;
perforation plates were simple (Fig. 14), transverse to oblique;
some vessels had 3 perforation plates; intermediate forms between
scalariform with one incomplete bar to irregular and reticulate
were observed (Fig. 15). Parenchyma was scanty paratracheal (Fig.
12). Multiseriate wide and high heterogeneous rays (Fig. 13) were
composed of procumbent, upright or/and square cells, with lignified
and unlignified portions; starch grains occurred in ray cells
(Figs. 10, 12). Libriform fibres were present, rare septate. Bands
of unlignified parenchyma including axial and radial parenchyma
were present in fibrous wood (Fig. 10). Root. Adult roots were
covered by a peridermis, similar to the stem (Fig. 16); composed of
pheloderm formed by 5 to 6 layers of rectangular cells in cross
section, with primary walls; cork was formed by several layers of
cells with suberized walls, which alternate with layers of cells
with lignified walls.
Cortex was composed of parenchyma cells with primary walls.
Druses were less frequent in E. calochlora, and absent in E.
rhodotricha .
The primary phloem was collapsed (Figs. 16, 17), with caps of
sclereids outside secondary phloem (Fig. 17). Secondary phloem
(Fig. 17) comprised narrow and short sieve tube elements, with
simple sieve plate, transversal to slightly oblique; companion
cells; axial and radial parenchyma. Secondary xylem was fibrous in
both species (Figs. 16-20), diffuse porous, composed of vessels
predominantly solitary and multiples of 2 to 3, and less frequent,
of 4 to 5 vessel elements with simple perforation plate, transverse
to oblique; in E. rhodotricha, intermediate forms between
scalariform with one incomplete bar to irregular and reticulate
were observed, similar to the stem fibrous wood; intervessel pits
were bordered; vessel-parenchyma pits were simple to half-bordered,
scalariform, opposite to alternate. Wide band tracheids were
absent. Scanty paratracheal parenchyma present (Fig. 17).
Unlignified, multiseriate wide and high heterogeneous rays (Figs.
16-19) were composed of procumbent, upright or/and square cells.
Druses and starch grains (Fig. 19) occurred in ray cells. Libriform
fibres were present, septate (rare) only in E. rhodotricha (Fig.
20). Unlignified parenchyma was present including also axial
parenchyma, forming bands, more evident in E. rhodotricha.
Discussion
The anatomical characteristics observed in the studied species
are commonly observed in other Cactaceae species, as extensively
described in the literature (Terrazas-Salgado and Mauseth, 2002;
Soffiatti and Angyalossy, 2003, 2007, 2009; Terrazas and Arias,
2003; Arruda et al., 2005; Mauseth, 2006).
The 2 Echinopsis species have a thick cuticle, an important
adaptation for xeric habitats, protecting the plant body from water
loss and pathogens. Unusual in Cactaceae (Gibson and Nobel, 1986;
Loza-Cornejo and Terrazas, 2003), they showed a secondarily
biseriated epidermis, which has also been reported for a few other
genera in Trichocereeae, such as Harrisia (Mauseth et al., 1998); a
multiseriate epidermis was mentioned for Espostoa Britton and Rose
(Mauseth, 1999). Reports of multiseriated epidermis exist for other
tribes such as Pachycereeae (Gibson and Horak, 1978; Barthlott and
Hunt, 1993; Terrazas-Salgado and Mauseth, 2002), Cacteae
(Terrazas-Salgado and Mauseth, 2002), Notocacteae (Nyffeler and
Eggli, 1997; Terrazas-Salgado and Mauseth, 2002), Cereeae (Darling,
1989; Mauseth, 1996), and Browningieae (Mauseth, 1996). As stated
by Loza-Cornejo and Terrazas (2003), multiple epidermis in cacti
occur in unrelated genera and seem to have appeared several times
in Cactaceae.
Parallelocytic stomata and a long substomatal chamber were
observed in the investigated species, a common feature for the
members of the family (Gasson, 1981; Eggli, 1984). In contrast with
most xerophytes, several species of Cactaceae show superficial
stomata (Eggli, 1984; Gibson and Horak, 1978; Fahn and Cutler,
1992; Loza-Conejo and Terrazas, 2003; Soffiatti and Angyalossy,
2007), where guard cells are on the same level as the other
epidermal cells. The long substomatal chamber, due to the thick
hypodermis, reduces water loss when the stomata are open (Fahn and
Cutler, 1992). According to Darling (1989), the reduced
transpiration rates are combined with low CO2 and O2 exchange
rates, prioritizing water conservation. Sunken stomata were
observed in E. rhodotricha, a situation reported for some other
genera (Metcalfe and Chalk, 1950; Gasson, 1981; Eggli, 1984;
Loza-Cornejo and Terrazas, 2003), maximizing water loss prevention.
Nevertheless, this is a variable feature in the genus, once
superficial stomata were observed in other species, such as in
Echinopsis aurea Britton et Rose (Eggli, 1984), and in E.
calochlora, while in E. eyriesii (Turpin) Zucc. these are sunken,
as in E. rhodotricha.
The organization of cortical tissues in both studied Echinopsis
species is typical of cacti (Gibson and Nobel, 1986; Loza-Cornejo
and Terrazas, 2003; Mauseth,
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10.7550/rmb.28124
Figures 13-20. 13-15, stem secondary xylem; tangential sections.
Echinopsis rhodotricha. 13, large rays. 14, simple perforation
plate of a vessel element. 15, incomplete perforation plate of a
vessel element. 16-20, root and cross sections (16-19). 20,
tangential section. 16-17, E. calochlora. 16, general view; note
peridermis composed of several layers of cells with lignified walls
alternate with cells with suberized walls; fibrous wood with large
unlignified rays. 17, phloem and xylem. Collapsed phloem (arrows)
outside functional secondary phloem. 18-19, E. rhodotricha. 18,
fibrous wood with large unlignified rays. 19, detail of unlignified
rays containing starch grains. 20, E. calochlora tangential
sections; septate fibres (arrows). Scale bars: 13= 400 m; 14-15= 10
m; 16, 18= 500 m; 17= 100 m; 19-20= 200 m. Co-cortex; P-
peridermis; Ph- phloem; R- rays; Xy- xylem
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1042 Garcia et al.- Anatomy of two species of Echinopsis
2006; Soffiatti and Angyalossy, 2007): there is a thick
hypodermis composed of several layers of colenchymatic cells,
followed by a photosynthetic palisade parenchyma and the inner
cortex, composed of isodiametric parenchymatic cells. The
hypodermis, being a flexible tissue, has important functions in
giving support to the stem and accommodating the stem changes in
volume due to shrinkage and swelling, also protecting the inner
tissues against pathogens (Gibson and Nobel, 1986). As the stems in
most cacti are the main photosynthesizing organ, the organization
of the chlorenchyma or palisade parenchyma is very similar to a
leaf (Sajeva and Mauseth, 1991). The internal cortex cells store
water (Mauseth, 1993b), and the large cells are able to expand and
contract depending on hydration state or water availability
(Mauseth, 2006).
Both species presented cortical and medullary bundles with
secondary growth, spread out in all directions very similar to leaf
veins. Like leaf veins, they are collateral and form a network that
extends to the base of the palisade parenchyma. Cortical bundles
are considered a synapomorphy for Cactoideae, present in nearly all
species (Mauseth, 2006). They perform important functions in sugar
and water transport to and from cortex cells and to the vascular
system (Mauseth and Sajeva, 1992; Terrazas-Salgado and Mauseth,
2002; Mauseth, 2006). Mauseth (2004) considers that the presence of
cortical bundles explains the large volume of cortex, as succulent
euphorbias, for instance, do not possess cortical bundles and their
stems never reach such a large size as cacti. Medullary bundles
have the same function as cortical bundles, transporting water and
nutrients (Mauseth, 1993a, 2006; Terrazas-Salgado and Mauseth,
2002), occurring only in Cactoideae (Mauseth, 1993a, 2006).
According to Gibson and Nobel (1986) pith diameter increased during
Cactaceae evolution and this event probably led to the emergence of
medullary bundles.
In the basal and older regions of the stem, as well as in the
root of these 2 Echinopsis species, the epidermis is substituted by
a peridermis, as noted for many species in the family. The origin
of phellogen from periclinal divisions of epidermal cells is very
common in cacti (Gibson and Nobel, 1986; Terrazas-Salgado and
Mauseth, 2002; Soffiatti and Angyalossy, 2003; Mauseth, 2006).
Although the origin of phellogen in a plant organ can be variable,
from different cell types, such as epidermal, colenchymatic,
parenchymatic or even phloematic cells, the most common origin is
from subepidermal layers of cells, according to Evert (2006). It
was seen in E. calochlora and E. rhodotricha that phellem in both
root and stem is composed by layers of suberized cells which
alternate with layers of lignified cells, also a common feature for
species of the family (Mauseth, 2006).
The phloem was composed of sieve tube elements and companion
cells, axial and radial parenchyma and sometimes, non functional
phloem might become lignified, similar to what was described for
several Cactaceae stems (Mauseth, 1999; Terrazas-Salgado and
Mauseth, 2002; Soffiatti and Angyalossy, 2003, Arruda et al., 2005;
Mauseth, 2006), and roots (Mauseth and Ross, 1988; Mauseth, 1989).
Echinopsis rhodotricha has fibre caps outside phloem; in E.
calochlora they are absent, typical of species which have non
fibrous wood (Mauseth et al., 1998).
Echinopsis calochlora and E. rhodotricha present fibrous wood in
the roots while in the stems they both have non fibrous wood,
corroborating the fact established by Mauseth and Stone-Palmquist
(2001). These authors stated that the structure of wood in the stem
and root of the same plant can be very different. Echinopsis
rhodotricha has also fibrous wood in the stem, characterizing what
is termed dimorphic wood: when young the plant produces one type of
wood and later on, the cambium produces another type (Mauseth and
Plemons, 1995; Mauseth and Plemons-Rodrguez, 1998). In the present
work, it was observed a typical situation in E. rhodotricha: which
first forms non fibrous wood, when the body is young and small and
does not need special investment in support, and when mature, the
cambium starts forming fibrous wood since support is then needed
due to the larger size (Terrazas-Salgado and Mauseth, 2002). The
fibrous and non fibrous wood types observed in the present study
have the same structure as described in literature, with fibrous
wood composed of vessel elements in a matrix of libriform fibres,
and non fibrous wood composed of vessel elements in a matrix of
wide band tracheids (Gibson, 1973; Gibson and Nobel, 1986;
Terrazas-Salgado and Mauseth, 2002; Mauseth, 2006; Soffiatti and
Angyalossy, 2009). The only difference between the fibrous wood of
the stem and root of E. rhodotricha is the presence of the storied
structure in the stem, absent in the root. A noticeable feature
observed in the secondary xylem of the stem in both species is the
occurrence of a storied structure in the secondary xylem. This
feature has been described in the literature for several species of
Cactoideae, including E. calochlora, by Gibson (1973). This author
observed that the rays are rarely storied, and mentions that the
storied structure is more evident in specialized growth forms, such
as globose and epiphytic species. Gibson (1973) also stated that in
non fibrous wood it is a difficult feature to distinguish due to
differences in the length of vessel elements and wide band
tracheids, but this was not the case in the present study. Storied
wood is a feature that occurs in several phylogenetically unrelated
groups and it is found in groups where fusiform cambial initials
are shorter (Carlquist, 2001). In the present study,
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10.7550/rmb.28124
the non fibrous wood showed a storied structure, especially
regarding wide band tracheids, more pronounced in E. calochlora
than E. rhodotricha.
Many of the characteristics presented in this work corroborate
the descriptions that had already been reported for other species
of Cactoideae. Some anatomical features can be used as diagnostic,
allowing for the segregation of the 2 species of Echinopsis
studied, such as: presence of biseriated epidermis, sunken stomata,
and fibrous and non fibrous wood in the stem of E. rhodotricha. As
Echinopsis comprises a large number of species (ca. 128 species,
Anderson, 2001), some intrageneric variability is expected for this
genus, and more species should be investigated, in order to
establish the diagnostic value of those traits mentioned.
Acknowledgements
Thanks to the Fundao de Apoio ao Desenvolvimento do Ensino,
Cincia e Tecnologia do Estado de Mato Grosso do Sul (FUNDECT), for
the master scholarship to the first author, to the Research
Coordinator Pro-Reitoria de Pesquisa e Ps-Graduao da Universidade
Federal Mato Grosso do Sul, from the Programa de Ps Graduao em
Biologia Vegetal, and the National Council of Research (CNPq),
process n. 473673/2007-0. We would like to thank the Scanning
Electronic Microscope Center of the Federal University of Paran,
especially to the Dr. Cleusa Bona due to the equipment use; to Dr.
Geraldo Alves Damasceno Jr. and the Msc. Vali Joana Pott for
identifying the species and to the Pantanal Seeds Net by the
logistic support on the field sampling.
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